The Early Phyllosoma Stages of Spiny Lobster Panulirus Echinatus Smith, 1869
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The early phyllosoma stages of spiny lobster Panulirus echinatus Smith, 1869 (Decapoda: Palinuridae) reared in the laboratory Abrunhosa, FA.a*, Santiago, AP.b* and Abrunhosa, JP.a aNúcleo de Estudos Costeiros, Campus de Bragança, Universidade Federal do Pará – UFPA, Alameda Leandro Ribeiro, Bairro Aldeia, CEP 68600-000, Bragança, PA, Brazil bLaboratório de Recursos Aquáticos, Universidade Federal do Ceará – UFC, Av. Mister Hull, s/n, Pici, CEP 60455-760, Fortaleza, CE, Brazil *e-mail: [email protected], [email protected] Received April 6, 2006 – Accepted September 1, 2006 – Distributed February 29, 2008 (With 6 figures) Abstract The early stages of the Panulirus echinatus were hatched and reared in the laboratory. Ovigerous females were cap- tured in their habitat and carefully transported to the laboratory. Larvae were transferred in a recirculation water tank at a density of 10 larvae.L–1. The larvae were fed on Artemia and gonads of mussel Brachydonts sp. Microalgae Dunaliella viridis was added at a concentration of 150 x 104 cell.mL–1. Larvae and exuviae of each zoeal stage were preserved in an alcohol 70% + glycerin (1:1) solution. The phyllosomas moulted eight times; the intermoulting period of each instar averaged about 7 to 10 days. The main morphological changes of each appendage were described in detail, illustrated and compared with previous reports. Keywords: spiny lobster, phyllosoma, larval development, morphology. Os primeiros estágios de filosoma da lagosta Panulirus echinatus (Decapoda: Palinuridae) cultivados em laboratório Resumo Os estágios iniciais de lagosta tropical Panulirus echinatus eclodiram e foram cultivados em laboratório. Fêmeas ovígeras foram capturadas em seu habitat e cuidadosamente transportadas para o laboratório. As larvas foram cultiva- das em tanques de recirculação de água e alimentadas com náuplios de Artemia e gônadas mexilhão Brachydonts sp. Microalga Dunaliella viridis foi adicionada em cada tanque de cultivo na concentração de 150 x 104 cel.mL–1. Os filo- somas mudaram oito vezes e o período de intermuda de cada instar foi de cerca de 7 a 10 dias. As principais mudanças morfológicas de cada apêndice foram descritas, ilustradas e comparadas com trabalhos anteriores. Palavras-chave: lagosta, filosoma, desenvolvimento larval, morfologia. 1. Introduction The Brazilian spiny lobsters of the Panulirus genus lected in the sea (Lewis, 1951; Baisre, 1964; Phillips and have high economic value. The species P. argus (Latreille, Sastry, 1980) and for those reared in the laboratory pro- 1804) and P. laevicauda (Latreille, 1817) are captured gressed through the metamorphic puerulus stage (Inoue, extensively, representing one of the most important fish- 1981; Kittaka and Kimura, 1989; Yamakawa et al., 1989; ery resources in the world (Phillips et al., 1980). For a Matsuda and Yamakawa, 2000). The long larval period long time another species, P. echinatus, was occasionally (about one year) is considered an important obstacle for captured, however, an increase in their production has phyllosoma culture research. The lack of detailed morpho- been observed (Santiago, 2001). Despite the economic logical description of some or all phyllosoma stages has significance that this latter species has represented in the also hindered studies of larval distribution and ecology. fisheries, few studies concerning their biological aspects Successful studies on culture from egg to the pueruli have been reported. stage of Panulirus were achieved for P. japonicus (Von Information on the biology of phyllosomas is consid- Siebold, 1824) (Kittaka and Kimura, 1989; Yamakawa ered important and necessary to elucidate aspects related et al., 1989) and P. longipes (A. Milne-Edwards, 1868) to larval biology and culture. The larval development (Matsuda and Yamakawa, 2000). Some attempts at rear- of spiny lobster has been reported for specimens col- ing phyllosoma larvae were reported for P. homarus Braz. J. Biol., 68(1): 179-186, 2008 179 Abrunhosa, FA., Santiago, AP. and Abrunhosa, JP. (Linnaeus, 1758) (Radhakrishnan and Vijayakumaran, gonads of Brachyodonts sp. Microalgae Dunaliella 1995) and for P. ornatus (Fabricius, 1776) (Duggan viridis Teodoresco 1906 were added daily in the culture and McKinnon, 2003) in which, 10 and 9 early instars system. were reared and described, respectively. More recently, Samples of larvae were removed after each instar Abrunhosa et al. (2004) described and compared the and fixed in 10% formalin solution for 24 hours and morphological characteristics of three early instars of P. preserved in a glycerol+alcohol 70% (1:1) solution. The laevicauda and P. echinatus, contributing to the identifi- larvae were dissected under an optical binocular micro- cation of phyllosomas that occur on the Brazilian coast. scope. The illustrations of each larval instar, including These three early larval instars of P. echinatus are again appendages, were made using a camera lucida. reviewed in the present study. For the species P. homarus, information on the av- The present study describes in detail and illustrates erage length, number of setae in the apical segment of the 5 early stages and instars of each stage of the spe- the 2nd maxilla and number of the exopod setae of each cies P. echinatus in the laboratory. Morphological com- appendage were based on the illustrations reported by parisons with larvae of its congeneric species, previously Radhakrishnan and Vijayakumaran (1995). described, are briefly discussed. The terminologies STAGES and INSTAR were used in order to make comparisons with other previously de- 2. Material and Methods scribed species. Both terms were extensively used by Ovigerous females of P. echinatus were obtained at the authors in descriptions of larval development of the Iparana Beach (Caucaia/CE) in October 2001 by autono- spiny lobster. mous diving in the coastal shallow waters. They were transported to the laboratory in plastic baskets (10 L) 3. Results containing local seawater. In the laboratory the females were acclimated and 3.1. On the larval culture then placed in a circular tank (cap. 1000 L) with biologi- The larval hatching was closely related to the moon cal filter and constant aeration. After hatching, the phyllosomas were carefully phases, in which the majority of the larval hatching was transferred to three transparent water recirculation tanks carried out on the first quarter and full moon, indicating at a density of 60 larvae.L–1. These tanks are similar to that lunar cycle had direct influence in the life cycle in those used for culture of many species of spiny lobsters the laboratory. of the family Palinuridae (Kittaka, 1994). The tanks, The phyllosomas readily feed on Artemia nauplii which were confectioned with acrylic (70 cm Ø and in stages I to III. Then, the phyllosomas seemed to 30 cm deeper), allow free circulation of flowing water have difficulty in catching Artemia so other foods were and total visualization of the floating elements is pos- tested. Some foods were initially well accepted by the sible. Salinity was maintained at 35‰. The larvae were phyllosomas, but later were rejected. This behaviour fed Artemia nauplii for the 3 first instars and then, with may be related to nutritional aspects, because many Table 1. Intermoulting period, accumulative days and maim characteristics of each stage for phyllosomas of Panulirus echinatus reared in the laboratory. Stage Intermoult Accumulative Main characteristics (Mean days) days I 9 9 Ocular peduncle unsegmented. Exopod of the 1st and 2nd pereiopods with 5 pairs of setae. II 9 18 Eyes segmented; exopod of the 1st and 2nd pereiopods with 6 pairs of setae. IIIa 7 25 Fourth pereiopod present; exopod of the 1st and 2nd pereiopods with 7 pairs of setae. IIIb 5 30 1st and 2nd pereiopods with 8 pairs of setae. IVa 5 35 Fifth pereiopod present; exopod of the 1st and 2nd pereiopods with 9 pairs of setae. IVb 5 40 exopod of the 1st and 2nd pereiopods with 10 pairs of setae. IVc 7 47 exopod of the 1st and 2nd pereiopods with 11 pairs of setae. IVd 8 55 exopod of the 1st and 2nd pereiopods with 12 pairs of setae. V >8* >63 Antennule 3-segmented; exopod of the 1st and 1st and 2nd pereiopods with 13 pairs of setae. *No phyllosoma succeeded in moulting to the next instar. 180 Braz. J. Biol., 68(1): 179-186, 2008 The early phyllosoma stages of spiny lobster phyllosomas failed to moult to the subsequent stage. This First Maxilla (Figure 2c): Coxal endite with 2 long fact was more evident for larvae in the VIII instar. serrulate setae and 1 short simple seta; basal endite with The larvae moulted eight times, once at stages I to II, 2 strong serrulate setae distally and 1-2 sub-distally; en- two times at stage III and four times at stage IV (Table 1). dopod short, unsegmented with 2 setae. There was not a specific day or night period in which Second Maxilla (Figure 2d): Basal segment en- the larvae moulted to the next instar. The moult of the larged with 2-3 short setae, distal segment short with phyllosomas from stage I to II and II to III ranged from 7 4 long plumose setae. to 11 and 7 days, averaging in 9 and 7 days, respectively. First Maxilliped: Small, uniramous and unsegment- For successive instars, it was not possible to determine ed protuberance. accurately the intermoult period for later instars, because Second Maxilliped (Figure 2e): Endopod 5-seg- the phyllosomas were not reared individually. The inter- mented, distal segment ending in 1 strong distal and moult average for each instar is shown in Table 1. 2 sub-distal spines, sub-distal segment rounded distally The largest mortality of the phyllosomas was on the with long serrulate setae; exopod absent. th 4 day of rearing, when about 50% of the larvae died in Third Maxilliped: Endopod well developed, 5-seg- both systems. After this period, low mortality rates were mented, segment distal with long setae rounding the observed.